Niche construction mediates climate effects on recovery of tundra heathlands after extreme event
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with pellets per year out of 144 plots, and belonged to reindeer (Rangifer tarandus tarandus), small rodents such as Norwegian lemming (Lemmus lemmus), ptarmigan (Lagopus lagopus), grey-sided mountain hare (Lepus timidus) and in the island of Rebbenes (Fig 1) the greylag goose (Anser anser). Experimental design We used a one-factor ANOVA design with extreme event (i.e. removal of all above ground vegetation) as the experimental treatment at each site (Fig 1). Removal of all above ground veg- etation was meant to imitate heavy interacting extreme events such as winter warming epi- sodes [7], burrowing by lemmings/voles [27, 28], or pathogens attacks [29] where large parts (if not all) of the aboveground vegetation dies or might be removed. Because certain types of extreme events can leave a litter layer (for example, a repeated winter warming event will kill the vegetation but not remove it, while voles cut the vegetation at ground level and remove it), this treatment had two sub-treatments: the cut vegetation was left as a litter layer or the cut vegetation was discarded, leaving a bare plot. In June 2009, ten blocks were established in each of the sites (Fig 1). Blocks were placed a minimum of five meters apart to avoid including the same E. nigrum individual in different blocks, since long-lived, clonally reproducing species can have multiple modular units [30]. Unfortunately, two blocks were removed from the study due to unexpected field work prob- lems (one at Skibotn and one at Guhkesja´vri, locations 4 and 5 respectively, Fig 1), leaving 48
blocks and 144 plots in total. Each block included three treatment plots of 50 cm x 50 cm each, i.e. control, extreme event without litter and extreme event with litter (Fig 1). The two extreme event plots were assigned treatments in a random manner and placed beside each other with a 50-cm separation and approximately 2 meters downslope of the control plot (Fig 1). Vegeta- tion was cut with standard garden shears at ground level including the vegetation in a 20-cm buffer zone between plots. Furthermore, all plots (except control plots) were trenched around with a spade which was 30 cm long to eliminate any below-ground effects of the vegetation adjacent to the plot. Environmental measurements Three temperature loggers (Thermochron iButtons1) were placed at ground level at each site to measure soil surface temperature every 3 hours all year round (during 2010–2019). This Table 1. Characteristics of the study sites. Rebbenes Skogsfjord Snarby Skibotn Guhkesja´vri Longitude/Latitude 70˚ 10’; 18˚45’ 69˚ 57’; 19˚ 15’ 69˚ 45’; 19˚ 30’ 69˚ 14’; 20˚ 33’ 69˚ 10’; 20˚ 42’ Elevation (m a.s.l.) 10 264 266 580 510 Mean annual temp. (˚C) 2.2 0.5 0.6 -2 -1.9 Mean annual prec. (mm) 871.18 860.49 917.19 634.16 628.2 Mean summer temp. (˚C, June-July-August) 12.37 11.77 12.04 11.96 11.64 Mean summer prec. (mm, June-July-August) 218.75 179.86 172.16 189.75 202.98 Continentality index 21.8 24.2 23 30.1 31 Freezing days (mean/year) 39 30 5 125 153 Soil pH 4 4 4.5 3.5 4 Soil organic horizon depth (cm) 5–10 3–10 2–5 3–10 5–10
Climatic variables were calculated based on data from the six study years using on-site soil surface temperature loggers, hence temperatures are measured at vegetation level. Precipitation data was gathered from publicly available databases (i.e. senorge.no). https://doi.org/10.1371/journal.pone.0245929.t001 PLOS ONE Niche construction effects on tundra after extreme event PLOS ONE | https://doi.org/10.1371/journal.pone.0245929 February 4, 2021 4 / 14